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Review Article
Abstract
Literature in evolutionary psychology suggests that mate choice has been the primary mechanism of sexual selection in humans, but this
conclusion conforms neither to theoretical predictions nor available evidence. Contests override other mechanisms of sexual selection; that is,
when individuals can exclude their competitors by force or threat of force, mate choice, sperm competition, and other mechanisms are
impossible. Mates are easier to monopolize in two dimensional mating environments, such as land, than in three-dimensional environments,
such as air, water, and trees. Thus, two-dimensional mating environments may tend to favor the evolution of contests. The twodimensionality of the human mating environment, along with phylogeny, the spatial and temporal clustering of mates and competitors, and
anatomical considerations, predict that contest competition should have been the primary mechanism of sexual selection in men. A functional
analysis supports this prediction. Men's traits are better designed for contest competition than for other sexual selection mechanisms; size,
muscularity, strength, aggression, and the manufacture and use of weapons probably helped ancestral males win contests directly, and deep
voices and facial hair signal dominance more effectively than they increase attractiveness. However, male monopolization of females was
imperfect, and female mate choice, sperm competition, and sexual coercion also likely shaped men's traits. In contrast, male mate choice was
probably central in women's mating competition because ancestral females could not constrain the choices of larger and more aggressive
males through force, and attractive women could obtain greater male investment. Neotenous female features and body fat deposition on the
breasts and hips appear to have been shaped by male mate choice.
2010 Elsevier Inc. All rights reserved.
Keywords: Evolutionary psychology; Contest competition; Mate choice; Mating; Sexual selection
1. Introduction
Viewing human mating in a developed nation, one
surmises that success in heterosexual competition for mates
entails attracting members of the opposite sex. Beauty,
fashion, and physical fitness are so important in places like
the United States that they have become multi-billion dollar
industries. Men and women have virtual autonomy to choose
their mates. These conditions are so pervasive that it is
tempting to think that they have characterized our evolution
that humans evolved in a context where, in the mating
arena, the preferences of the opposite sex were the primary
forces shaping our phenotypes.
With notable exceptions (e.g., Apostolou, 2007; Archer,
2009; Buss & Dedden, 1990; Buss & Duntley, 2006; Buss
& Shackelford 1997; Daly & Wilson, 1988; Lassek &
Gaulin, 2009; Sell et al., 2009), the recent literature in
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2. Sexual selection
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Dimensionality of mating
environment
Body size
Capability of physically
constraining opposite sex
Temporal clumping of
available mates
Spatial clumping of mates
and competitors
Females
2D (Contests)
2D (Contests)
Large (Contests)
Yes (Contests)
Large (Contests)
No (No Contests)
No (Contests)
No (Contests)
Multi-male/
multi-female groups
(Contests reduced)
No (Contests)
Multi-male/
multi-female groups
(Contests reduced)
No (Contests)
Body-size constraints of
flight or arboreality
Contests in close phylogenetic Yes (Contests)
relatives
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No (No contests)
Thompson, & Wrangham, 2007; Nishida & HiraiwaHasegawa, 1987; Rodman & Mitani, 1987; Smuts, 1987).
Only multi-male groups are predicted to reduce individual
males' abilities to monopolize females, elevating the
importance of other forms of sexual selection, but the
influence of this variable may be mitigated in humans
(see below).
Female contests are absent in humans' close phylogenetic
relatives, and monopolization of mates is likely to be
unfeasible in women, as well (Table 1). The multi-female
structure of human groups should have hindered ancestral
females from excluding their competitors from mates.
Indeed, in the presence of multiple same-sex competitors,
successful mate defense probably depends partly on mates'
cooperation. However, evolutionary models suggest that
such cooperation will tend to evolve in one sex when the
other is physically dominant and thus capable of sexual
coercion (Clutton-Brock & Parker, 1995). Yet, men are
greater in size and physical prowess than women are, and
thus, men should be prohibitively difficult to constrain in
their choices. We can therefore predict that female mating
competition would favor traits to attract men, rather than
physically monopolize them.
Before these predictions can be evaluated, it is necessary
to clarify what constitutes evidence of a trait's adaptive
function in winning mates. For example, is a particular trait a
weapon or an ornament, a dominance signal or a mate
attraction display?
3.1. Testing evolutionary predictions
One can infer ancestral selection pressures by studying
the adaptations that they produced. Natural selection is the
only evolutionary process to systematically produce traits
that appear engineered for specific functions (Williams,
1966). If, under scrutiny, a trait looks well-suited to a
purpose that would have benefited ancestral bearers, then we
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in correlates positively with peers' rankings of his dominance (Pellegrini, 1995; Pellegrini & Smith, 1998). Men
report engaging in, and inclinations to engage in, nearly one
standard deviation more physical aggression than women
(Buss & Perry, 1992). Men perpetrate more offensive
physical aggression, defined as non-defensive attacking,
hitting, and/or restraining another individual in all societies
studied (Ellis et al., 2008). The vast majority of same-sex
homicides (about 95%), from every society and time period
for which data are available, are committed by men (Daly &
Wilson, 1988; M. Wilson & Daly, 1985). Importantly, these
data do not include war killings, which occur almost entirely
at the hands of men (Adams, 1983). Traumatic injuries in
ancient skeletal remains indicate that interpersonal violence
was especially prevalent among men throughout human
history and prehistory (Walker, 2001).
Such sex differences in traumatic skeletal injuries may
help explain why some aspects of men's skeletons,
particularly in the face, are more robust. For example, in
modern populations, the incidence of mandibular fractures is
approximately five times higher in men than in women,
young men are disproportionately represented, and the
primary cause is typically found to be violent assault with
a fist or blunt object (Adi, Ogden, & Chisholm, 1990; Haug,
Prather, & Indresano, 1990; Scherer, Sullivan, Smith,
Phillips, & Robson, 1989; Sojat, Meisami, Sandor, &
Clokie, 2001). A similar pattern in the evolutionary past
could have selected for more robust mandibles in men than
in women.
Certainly, size, strength, speed, and aggression in men
correlate with physical competitive ability, and manipulations that increase these variables lead to greater physical
prowess. This is why many athletes abuse anabolic steroids.
Relatively greater male upper-body (compared with lowerbody) muscle mass and strength in particular suggest an
evolutionary history of fighting (Sell et al., 2009). These
traits also characterize male contests across species; males
are larger, stronger and more aggressive in diverse species
with male contests across the animal kingdom (Andersson,
1994). Close relatives of humans with minimal male
contests, such as gibbons, lack substantial sex differences
in size, strength, and aggression.
Men possess several traits that appear to function
primarily in threatening rivals. For example, beards and
eyebrow hair grow at puberty in males and may signal
dominance through association with testosterone levels and
by increasing the apparent size of the jaw and brow (Guthrie,
1970; Muscarella & Cunningham, 1996; Neave & Shields,
2008). Male faces with beards are rated as more dominant
than the same faces clean-shaven (Muscarella & Cunningham, 1996; Neave & Shields, 2008). Likewise, deep, lowpitched voices increase men's apparent size (Feinberg,
Jones, Little, Burt, & Perrett, 2005) and dominance (Puts,
Gaulin, & Verdolini, 2006; Puts, Hodges, Cardenas, &
Gaulin, 2007). Perhaps deep voices signal dominance in men
partly because they correlate with high testosterone levels
Fig. 3. From a great ape progenitor with single-male polygyny and singlemale social groups (1) (Harrison & Chivers, 2007), two trajectories for
African apes are depicted: a continuation of this pattern in Gorilla, and the
evolution of multi-male groups for female defense, as in African lions, in the
common ancestor of Pan and Homo (2). In Pan, within-group monopolization of females was difficult, and greater sperm competition thus
predominated (3), resulting in lower sexual dimorphism (Jungers & Susman,
1984) but larger investments in testicular tissue (Short, 1979). In Homo and
their immediate ancestors, individual males more effectively monopolized
females, perhaps due female cooperation, between-group competition, and
more intensive use of weapons.
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Fig. 4. Masculinity in facial hair, voice, facial structure and body build has
larger positive effects on perceptions of dominance than on perceptions of
attractiveness. Note: the important comparisons are the effects of
masculinity on attractiveness versus dominance within each study.
Between-studies comparisons are confounded by differences in the
magnitude of manipulations and other methodological details.
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4. Summary
The ancestral human mating system may have comprised
groups of (often related) males cooperating in female
defense. Between-group aggression, female cooperation,
and the ability to inflict lethal injuries with weapons likely
enabled some males to monopolize multiple females. At the
same time, female defense was imperfect, promoting
moderate sperm competition and female choice of both
long-term mates and extra-pair sex partners. Monopolization
of females was probably related to social skills and
attractiveness to females, but force or threat of force seems
to have been especially important. Moderate paternity
confidence coupled with efficient hunting promoted male
investment, which may have been elaborated as an
alternative mating tactic. Females may have evolved sexual
ornaments such as neotenic faces, high-pitched voices, and
fatty breasts and hips to attract male investment.
5. Conclusions
Human mating is complicated. It is the stuff of operas and
soap operas, full of manipulation and deception, aggression
and solicitude, cooperation and selfishness. It is the
culmination of multiple individual interests, sometimes
overlapping, often opposing. Human mating is perhaps
even more complicated than it appears in contemporary
industrial societies, where men and women choose their
mates largely beyond the authority of kin, women do not rely
economically on men, and men are prohibited by the state
from using force against mates and sexual competitors. As
complicated as human mating is, it is becoming clear that
contests must have been very important in determining men's
reproductive success, and male mate choice must have been
very important in determining women's reproductive success. We can predict this theoretically from the dimensionality of our mating environment, the structure of human
groups, differential parental investment, and phylogeny. We
can also see it in the traits that selection has produced.
The idea that male mate choice has been an important
selection pressure on women is relatively uncontroversial,
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